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A Revision of Amansia glomerata C. Agardh, Amansia rhodantha (Harvey) J. Agardh and Melanamansia glomerata (C. Agardh) R. E. Norris (Rhodophyta: Rhodomelaceae)
A. D. R. N’Yeurt
Jeune Equipe Terre Océan, Université de Polynésie Française, BP 6570, Faa’a Aéroport, Tahiti 98702, French Polynesia, firstname.lastname@example.org
Species of Amansia and Melanamansia from the tropical Pacific region have been re-examined, and compared with material from the Indian Ocean, Japan, Taiwan and the Philippines. All specimens examined from the Pacific region (except Hawaii and some from New Caledonia and Fiji) lack pseudo-pericentral cells, thus placing them in Amansia. Melanamansia glomerata is only reported from Hawaii, New Caledonia, Fiji, the Philippines,Vietnam, Malaysia and for the first time in the Indian Ocean from Kenya.Type material of Amansia rhodantha (Harvey) J. Agardh from Mauritius was compared and found not significantly different from the Pacific Amansia material (including type and isotype material of Amansia paloloensis South et Skelton from Samoa, which is reduced to synonymy), thus extending the distribution range of A. rhodantha as a pantropical entity.
The genus Amansia was erected by Lamouroux (1809: 322) and is placed in the tribe Amansieae of the family Rhodomelaceae (Hommersand 1963: 335). The tribe is characterised by dorsi-ventral, erect thalli arising in tufts from a common basal disc. Most genera have five pericentral cells around the axial cell (Figs 1, 2) except Kuetzingia Sonder and Protokuetzingia Falkenberg, which have six pericentral cells (Kylin 1956). The two first-formed pericentral cells lie on the dorsal side of the characteristically inrolled apex of the mature lanceolate blades. In Amansia the dorsal and dorsi-lateral pericentral cells further divide to form immediately behind the apex a distromatic wing or ala, whose cells are arranged in alternate layers, with secondary lateral pit-connections (Fritsch 1945: 570). Recently, Norris (1988b, 1995) re-examined species of Amansia in the Indian Ocean and Hawaii, and discovered that some members of this genus differed consistently by having the first and second pericentral cells each producing a lateral pseudo-pericentral cell lying adjacent, but not connected to, the axial cell (Norris 1988b: 214; Fig. 3). Species previously ascribed to Amansia that had these pseudo-pericentral cells were transferred to a new genus (Melanamansia R. E. Norris 1988b: 217). Notably, Hawaiian plants previously ascribed to the common tropical species Amansia glomerata C. Agardh (whose type locality is Hawaii) were transferred to Melanamansia glomerata (C. Agardh) R. E. Norris, while Indian Ocean ‘Amansia glomerata’ were put under the earlier name Amansia rhodantha (Harvey) J. Agardh based
on their lack of pseudo-pericentral cells (Norris 1995). The genus Amansia currently consists of four species, the Caribbean A. multifida Lamouroux (the type of the generic name), A. loriformis R. E. Norris and A. rhodantha (Harvey) J. Agardh from the Indian Ocean and Malaysia, and A. paloloensis South et Skelton from Samoa. Melanamansia currently consists of twelve species, M. daemelii (Sonder) R. E. Norris, M. glomerata (C. Agardh) R. E. Norris, M. japonica (Holmes) R. E. Norris, M. mamillaris (Lamouroux ex C. Agardh) R. E. Norris, M. pinnatifida (Harvey) R. E. Norris, M. pumila (Sonder) R. E. Norris, M. scalpellata (Tanaka) R. E. Norris, M. seagriefii R. E. Norris, M. serrata (Harvey) R. E. Norris and M. mitsuii (Tanaka) Yoshida. Following these revisions by R.E.Norris,several authors (Yoshida et al. 1995,N’Yeurt et al. 1996,Payri and N’Yeurt 1997, Millar 1999, Millar et al. 1999) made the nomenclatural change from Amansia glomerata to Melanamansia glomerata in their checklists and herbarium collections from the tropical Pacific, but until recently no detailed studies verifying these changes as correct have been done. In 1995, samples of ‘Amansia glomerata’ from Rotuma Island (north of Fiji) were sent to Dr R.E.Norris,who confirmed (pers. com.) that pseudo-pericentral cells were present, and on the basis of this assumed finding the name Melanamansia glomerata was used in later publications (N’Yeurt 1996,N’Yeurt et al. 1996).Later,in the course of on-going investigations on the Samoan algal flora, South and Skelton (1999) found a species of Amansia lacking pseudo-pericentral cells, which they described as new (A. paloloensis South et Skelton), while in Malaysia and Vietnam (Masuda et al. 2000, Masuda
Macrophotography was done with a Nikon F2A in conjunction with Kodak Plus-X pan 125 film. Palolo Deep. leg. Sakai and S. 6 Jun. N’Yeurt (UPF 347). Results Representative material examined Amansia rhodantha – Japan: Hanasado Beach. deep algal rim. The results of these investigations are described below. – Cook Islands: Ngatangiia. SUVA-A 5436L). SUVA-A 1566 (isotype). 1999. Fig. P. otherwise as in Holmgren et al. P. 5481L (spermatangial)]. A. Fig. SUVA-A 5438L). A. developed and printed in the laboratory. leg. M. Figs 1–3. 19 Mar. Kagoshima: 9 August 1959. – Taiwan: Anon. N’Yeurt In order to clarify the taxonomy of species hitherto ascribed to Melanamansia glomerata. 3896. Skelton. Okinoerabujima Island (60 km north of Okinawa). Tahiti. Material and Methods Specimens were collected by reef-walking. Prepared slides were examined using a Zeiss compound microscope. Okidomari. Tanegashima Island. 3898. Fig. 1998. 25 October 1984 (SAP 046978). in EPHE).Agardh to co-exist in the flora. Tahiti. Arrangement of pericentral cells in species of Amansia and Melanamansia (modified from Norris 1988b. 3892. Afiti and D. pers. Tombleson and T. Fiji. New Caledonia. 3895. 1998. M. A. D.‘Al’ and ‘CG’ refer to specimens housed at IRD. 13 May 1982. – French Polynesia: Moorea. Photomicrographs were obtained using an Olympus BX-50 compound microscope. Nouméa. 1998. showing intrusion of the first derivative of the first-formed pericentral cell into the axial ring (Suva. 1984. Rarotonga. A. Amansia rhodantha (Beqa. Skelton and Dave (SUVA-A 1567). Most recently. A. N’Yeurt (UPF 584). snorkelling or SCUBA diving. 3893. . Skelton. leg. 1998. 3899). Sections were made using a freezing microtome. M. leg. 1. 2. leg. Taharaa. in particular paying attention to important anatomical features such as the presence or absence of pseudo-pericentral cells and the number of segments between lateral veins of the blade (Norris 1995). A. P. U. E. 18 Sep. leg. 1997. Y. – Samoa: Palolo Deep Reserve. 9 Feb. 3897). SUVA-A 2402 (Lectotype) and liquid-preserved isotypes 5479L (tetrasporic). 11 Jul. Enomoto (SAP). Rintels (SUVA-A 3891. 3. leg. R.. Skelton and Dave [SUVA-A 1565 (holotype). Payri (8/84. leg. Melanamansia glomerata. 2 Jul.232 A. and stored in 5% buffered Formalin in seawater prior to shipment to the laboratory. 1–5 = order of formation of pericentral cells. 1989. Faletose (SUVA-A 3894. A. and stained with cotton blue in lactophenol or 1% acidified aniline blue in 60% clear corn syrup. Amansia rhodantha. Amansia rhodantha and Amansia glomerata. C. N’Yeurt (SUVA-A 5128). and a camera-lucida attachment was used to make anatomical drawings. Baba (SAP 056163). Baba (SAP 056160). P. 1998. leg. Okinoerabujima Island (60 km north of Okinawa) 27 Sept. ps = pseudo-pericentral cells). Fiji.) recent studies found both Melanamansia glomerata and Amansia rhodantha (Harvey) J. it was decided to revise records in the light of recent findings. (1981). a detailed morphological and taxonomic study on some selected members of the Amansieae (Wilson and Kraft 2000) echoed the opinion that the genus Amansia in Australia was in need of critical revision. 5480L (cystocarpic). 2 Jul. com. Furusato. leg. Punaauia. a = axial cell. leg. cross section of the lectotype from the Hawaiian Islands (LD 42600) showing the first two dorsal pericentral cells each developing a pseudo-pericentral cell (ps) that lies adjacent to the axial cell.
1067). leg. R. leg. Hildebrandt (L 0108596. leg. E. Cordero and Tuangpalan (US 091395).. Sumbawa. 1995. P. Barrow Island.Pagodpud. leg. leg. Kasahara (SUVA-A 1012.5 3. leg. leg. Daida (US 086755). Salayer. A. Huisman. 1995. Huisman (MURU. Amansia rhodantha and Melanamansia glomerata surveyed in this study. 84. 1876. A.. 18 March 1992. B79). 14 March 1998. P. Oahu. leg.Dumaguete City.Siquijor Is. JH240). L 0108598). J. A. 1985. Candelaria Zambales. 1876. Magruder (US 086459). M. n.leg. LD 42600 / 0772).2 53. – Fiji: ‘Feejee’. Suva Reef.Negros Oriental.Palawan.Ilocos Norte. 84 (L 0108603). Oahu. Potipot Is. 1965. leg. leg. Farlow (L 0108597). Bahati. Kauai.d. 24 April 1996. SIPHILEXP-78 (US 088531). 15 Oct. 26 Nov. A. Negros Oriental.. in TCD (Telfair. leg. leg. Kauai. 1983. Masuda et al. Yasawa Group. 1981. Garrigue (CH1). 1997. leg.9 18. Huisman (MURU. leg. 06 Sept. Hanauma Bay Beach Park. 1996: 472. paloloensis and Melanamansia glomerata specimens examined in this study. Carlson (SUVA-A 281).14 May 1979. 1069). 1984. Richer de Forges (AL 288d-CG 77a). SIPHILEXP-78 (US 013760). E. – New Caledonia: Lajeu Sudouest.d. n. c. W.23 May 1978. A.d. Basay. Suva Barrier Reef. leg..Revision of some Amansia and Melanamansia species 233 Table I. Agardh).7 2 2 1 2 2 2 2 2 2 * As measured on an index of (1 = little) to (3 = pronounced) – Rotuma: Hoféa. SIPHILEXP-78 (US 088532). J. leg. Lopta. Naukathura Island. 13 July 1972. Cannon (SAP 052062). 1982. Wilkes Exploring Expedition (US 04057b) (doubtful record. 1954. Bay of Sanggar. – Philippines: Actin. leg. D. 07 Oct. n. Ovalau. Jurien Bay WA. B. Average values of selected characters for Amansia paloloensis.d. – Lord Howe Island: 18 Aug. A. R. G.6 3. Lawai Kai. 28 Nov. W. 27 Dec. R. leg. Nanuyalevu. Silva et al. Norris 1988b: 211. M. Tantabiddi. 5 June 1967. Cribb (SAP 051714). A.. H. 16 Nov. leg. R127). leg.Canipo Is. Garrigue (CG 77c. leg. – Kenya: Mombassa. M de Robillard (L 0108599). Dumaguete. E. 1994 (SUVA-A 797). Banilad. Viti Levu. 1985.P. Amansia rhodantha: n. Lectotype. leg.d.7 54. Snellius-II (L 0108602). 14 Feb. 16 Oct. H. 6 Jun 1978. 84. leg.Calindagan Reef. fide G. Villeneuve (SUVA-A 1737). figs 1–11 (as Amansia glomerata C. 24 Nov. C. – Indonesia: Cape Batu Kerapo. 30 Jun. 1834).Moreland (US 50851). Gaudichaud (‘Amansia glomerata’. 1819. Melanamansia glomerata – Hawaiian Islands: Hawaiian Islands. – Mauritius: Delesseria rhodantha Harvey (holotype) (= Amansia rhodantha): Cap Malheureux. Taxonomic results Amansia rhodantha (Harvey) J. Agardh 1841: 26. Species Thallus height (cm) Blade width (mm) Blade length (mm) Serrations (1 –3)* Cortication (1 –3)* Rosettes (1 –3)* Amansia rhodantha Amansia paloloensis Melanamansia glomerata 39. leg. 1982. Garrigue (SUVAA 5436L). anon. B.3 2. 17 Mar.. 1979. H. Hildebrandt (L 0108600). Vatualailai. Isaac (L 0108595). Mauritius. Viti Levu. Ti Tree.Cuyo.Negros Oriental. Wilkes Exploring Expedition (US 04057a). A. C. 1995. G. 10 July 1992. – Kenya: Mombassa. 17 Jul. Tangangge. J. leg. leg. Table I presents the average results of morphological measurements carried out on Amansia rhodantha. leg. Ballou (SUVA-A 5438L). M.leg.Siaton.Solong-On. leg. leg. Snellius-II (L 0108601). 1979. SIPHILEXP-79 (US 088533). N’Yeurt (SUVA-A 346. S. – New Caledonia: Lajeu Sudouest. Oahu. AL 288b). Barrow Island. Isaac (L 0108594). leg. leg. Huisman (MURU. J. Magnesia Virae. – Philippines:Gao-oa. see text). SIPHILEXP-78 (US 088534). leg. Catanduanes. South (SUVA-A 5544.1 22. M. J. South in lit. leg. leg. 16 March 1991. C.). 1973. leg. G. J.9 11. 1980. leg. C. G. 1984. leg. leg. S. Unia. W. 10 May 1978. (US 02890). 29 May 1978. Cordero and Tuangpalan (US 091394). leg. – Mozambique: Ponta Abril. W. N’Yeurt (SUVA-A 5437L). Figs 4–10 .Malo. Ningaloo. 3 and 5 March 1992. Plongeurs IRD (AL 288a. Hapmak.. South (SUVA-A 5431). Eagles Nest. Negros Oriental. Villeneuve (SUVA-A 1774). – Australia: Heron Island QLD. Jul. 1838–42. leg. Diani Beach. leg. Chesterfield. Littler (US 163490). (MURU. leg. 1988. figs 37–42. CG 77b). R. leg. NR 248).11 Feb. n. 25 Feb. South (SUVA-A 2460). – Fiji: ‘Feejee’. 21 Sep.. Namada. 3 and 15 Oct. Montebello Islands. Viti Levu. J. Rukuruku Village. leg. Inhaca Peninsula. B262). Beqa Lagoon. Yasawa Islands. 3 Feb. 21/22-Sept. Skelton and G. 9 Sept. 10 Aug. Jul. Kipukai. 3 Jun 1978. leg. Keats. 1980. leg. Hakula. Kasahara (SUVA-A 1013. SIPHILEXP-78 (US 088530).W. R. P. Huisman (CWC 1997. 25 September 1999. B. 1838–42. 2000: 188. 347).
7. South Pacific (scale bar = 10 µm). 8. Type material of Delesseria rhodantha Harvey. and a pair of budding branchlets from the midrib in the lower half. from Rotuma Island. 5. 6. showing both lax and dense growth forms. Surface view of segment cells of blade alae. Fig. US 02890 (scale bars = 3 mm). 10. Fig. Habit of mature plant from the type locality. Detail of tetrasporangial stichidia on lateral endogenous branch.234 A. 9. Figs 4. Mauritius. R. Rehydrated type material of Delesseria rhodantha Harvey. Type material of Delesseria rhodantha Harvey. showing axial cell (a) surrounded by five pericentral cells. Secondary branchlet. Fig. SUVA-A 5437L. N’Yeurt Figs 4–10. D. in TCD (scale bar = 200 µm).5 mm). Fig. in TCD (scale bar = 2. Axial structure of blade. . Amansia rhodantha. Indian Ocean. in TCD (scale bar = 25 µm). showing marginal serrations.
Detail of the holotype (SUVA-A 1565). 13.Revision of some Amansia and Melanamansia species 235 Figs 11–16. SUVA-A 3892 (scale bar = 2. 12. Note marginal cystocarps (arrowheads). Habit of mature plant. Fig. Fig. 11. 4. . Detail of blades. Segment cells of blade alae in surface view. SUVA-A 5479L (scale bar = 1 mm). SUVA-A 3892 (scale bar = 12 mm). Fig. Holotype of Amansia paloloensis. SUVA-A 1565 (scale bar = 25 µm). Detail of morphologically distinct mature rosette and corticated axis.5 mm). 15. showing lateral endogenous branches and midrib. Amansia paloloensis from Samoa. 16. Tetrasporangial stichidia of Amansia rhodantha from Beqa. SUVA-A 5436L (scale bar = 50 µm). Fig. Fig. with numerous rosette axes. Compare with Fig. showing both lax branching and the presence of rosettes (arrowheads) (scale bar = 4 mm). Fig. 14. Fiji.
17. Norris 1995: 66) is on average 4. and two displaced rows of regularly arranged hexagonal alar cells 20–29 × 92–129 µm in surface view. Fig. XXV. Okamura 1900: 71. Axial structure of plant from Lajeu. 9). 19. fig. figs 14–29. Abbott 1999: 404. US 4057b (scale bar = 1 mm). pl. CG77a (scale bar = 10 µm). 31 fig. Individual tetrasporangia are spherical and tetrahedrally divided. 21. Vietnam. on average 53 mm high. 6E (unverified record). stem-like and rigid in basal portions. 6. each cell is about 20–20 × 87–100 µm in surface view. they are up to 4 cells long and non-vesiculate. Tropical Pacific Ocean. with a central axial cell 28–30 µm in diameter surrounded by five pericentral cells 36–40 µm in diameter (Figs 1. pl. 436B. figs 1–13. 20. 185. Jaasund 1976: 131. Fig.4. New Caledonia. or with regular serrations (endogenous branches). 22. Procarps and curved spermatangial heads are 200–450 µm in diameter and are dorsal on the involucral tips of endogenous lateral branches. E. fig. Lewis and Norris 1987: 22. colourless and uniseriate dorsal trichoblasts up to 8 cells long develop on every second or third axial cell (or on every segment in determinate marginal branchlets). Tetrasporangia occur in flattened. with a range of 1–5. 25–42 × 83–125 µm. Isaac and Isaac 1968: 24. Payri and Meinesz 1985: 511. with a range of 3–6. Silva et al. Distribution: Tropical Indian Ocean. pl. Fiji. Samoa). figs 2–14 (type locality: Palolo Deep Marine Reserve. and are initially vesiculate. 1996: 87. composed of two overlapping layers of elongate hexagonal cells in transverse rows. Kenya. The thallus is rose-red. 197. Tsuda and Wray 1977: 103. 1995: 150. Internal structure is cellular. – Melanamansia glomerata (C. New Caledonia. Blades of plant assumed to be from the Fiji Islands (see text). with in-rolled leaf apex. 1. Kenya. 267. N’Yeurt 1996: 428. The main axis is irregularly branched. Agardh) R. Payri and N’Yeurt 1997: 897. Kenya. Coppejans and Millar 2000: 333 (unverified record).5–0. Ultimate branchlets are ecorticate. Detail of blade of plant from Mombassa. surrounded by five pericentral cells.5 µm in diameter that lies adjacent to the axial cell (Figs 3. A distinct midrib up to 1 mm wide is present in the lower half of ultimate branchlets. 100–112 µm in diameter. 1. 128. Fig. Millar et al. New Caledonia. Deciduous. L 0108598 (scale bar = 5 mm). Figs 11–16 Misapplied names: Amansia glomerata C. 4B. . fig. to 0. Melanamansia glomerata (C. pl. Blade margins are smooth. e–i. Kylin 1956: 544. Japan.8–14. 18. N’Yeurt Basionym: Delesseria rhodantha Harvey 1834: 151–152. The number of segments between alternating veins (determinate branches covered by the corticating pericentral cell derivatives. Agardh 1824: 247. Norris 1988b: 211. L 0108598 (scale bar = 700 µm). Fritsch 1945: 570. figs 1–11. up to 750 µm long and 250 µm broad. up to 35 mm long and 6 mm broad. N’Yeurt et al. Tetrasporangial stichidia are pinnately arranged on either side of blade margins. in up to 10 regularly disposed pairs. Apia. Mauritius. Fig. Synonymy: Rytiphlaea rhodantha (Harvey) Decaisne 1842: 358. pinnately arranged marginal stichidia up to 340 µm long and 290 µm in diameter. Branching is lax to profuse and usually forms deep-red leafy rosettes of secondary branchlets when fully mature. showing smooth margins and elongate shape. Falkenberg 1901: 416. Millar 1999: 523 (unverified record). Plant from Potipot Island. 24. Trichoblasts are sparse. Norris 1995: 67. while cystocarps 700–1000 µm in diameter occur usually singly and terminally on endogenous branches. E. 7 mm broad. 2. Philippines. The central axial cell is 28–30 µm in diameter. Garrigue and Tsuda 1988: 63. 19). 77–110 µm thick. Fig. Blade of plant from Lajeu. 23. East Africa. The number of segments between alternating veins (determinate branches covered by the corticating pericentral cell derivatives. and are composed of lanceolate to ovate blades to 100 mm long and 3 mm wide. Fig. Figs 17–24 Basionym: Amansia glomerata C. Yamada and Tanaka 1938: 86. with the first two dorsal pericentral cells each developing a smaller pseudo-pericentral cell 13. Womersley and Bailey 1970: 336. AL288b (scale bar = 5 mm). becoming narrower and disappearing towards the apex. Tetrasporangial stichidia of plant from Oahu. showing pseudo-pericentral cells (arrowheads) surrounding (but not directly connected to) axial cell (a). Weber-van Bosse 1923: 369. 0. Amansia paloloensis South et Skelton 1999: 247. fig. The stem of plants is thick and cartilaginous. 1987: 61. containing up to 10 Figs 17–24. 198. Norris 1995: 67. 1999: 573. D. denuded below. 212a. Agardh 1824: 194. 85–110 µm thick. 20. figs 1–3 (type locality: Cap Malheureux. R. US 091395 (scale bar = 5 mm). holotype in TCD.8 mm in diameter. figs 127. Dublin). SUVA-A 5431 (scale bar = 50 µm). fig. Philippines. Note that the stichidia is twice laterally branched. Isaac 1956: 188. Known distribution: Hawaiian Islands. figs 19–21. and are crisp and lanceolate. Plants are brownish-red. Norris 1995: 66) is usually 4. South-East Asia. A central midrib is present. Yoshida et al.236 A. up to 100 (on average 40) mm high. with more or less pronounced marginal serrations 1–4 mm long (which represent endogenous branches) always present. New Caledonia. pl. CG77a (scale bar = 1 mm). Agardh) R. plate CXXVI. and the apex is characteristically in-rolled. Carposporangia are lanceolate. 118C–J. Fig. Melanamansia glomerata. Marginal teeth are up to 1 mm long and 445 mm broad. Plant from Unia. Fig. Secondary blades develop as successive pairs on the dorsal side of the midrib. Cribb 1983: 106. Plant from Mombassa. fig.
Spermatangial heads are curved and arranged in marginal clusters. In all cases.Revision of some Amansia and Melanamansia species 237 pairs of tetrahedrally-divided tetrasporangia which are 50–100 µm in diameter. the colour of the Melanamansia plants was noticeably brownish-red. procarps and cystocarps occur on endogenous marginal branches. . and stained the herbarium mounting paper.
rhodantha from the Indian Ocean and other localities. Exploring Expedition. MURU 248). while the other (US 04057b) is Melanamansia glomerata. length of the blades. a re-examination of the lectotype of Amansia glomerata C. as well as Amansia species from Taiwan. an additional field test to distinguish Amansia spp.). additional distinguishing characteristics of Melanamansia are a darkly coloured brown thallus which stains the mounting paper when dried (possibly linked to differences in chemical composition. rhodantha. 14) and corticated rosette axes (Fig. thicker. and absence of pseudopericentral cells. a comparison of material of A. more coriaceous branchlets with less apparent midribs (this study). which is welldocumented and illustrated in the protologue of the species. it appears to be conspecific with the latter species and thus extends the status of A. regularly alternate marginal serrations which are not as pronounced as in Amansia rhodantha and which are often absent. The status of Amansia paloloensis South et Skelton South and Skelton (1999) described a new species of Amansia from Samoa. 7) shows features similar to those seen in blades of South Pacific material previously ascribed to Amansia glomerata.S. the absence of blades organized in morphologically distinct tight axes around the parent axis. Melanamansia glomerata in the tropical Pacific Wilson and Kraft (2000: 366) commented that the generic segregation of Melanamansia from Amansia is probably not warranted. Amansia tends to bleach to translucent white. degree of cortication of the midrib and position of the reproductive stichidia are equally quite variable in plants from the same and similar populations. paloloensis from A. On the other hand. and trichoblasts lacking protective vesicles when young (Norris 1988b). rhodantha as a pan-tropical entity. Now that the Samoan species has been compared with genuine herbarium material of A. Masuda pers. plate CXXVI. The habit and description of the holotype by Harvey (1834: 151.238 A. when pseudo-pericentral cells were found to be absent) are considered conspecific with Amansia rhodantha. R. Kenya. In the Fiji Islands. a single record for this genus exists (collected from ‘Feejee’ by the Charles Wilkes U. M. material from the Tropical Pacific and Indian Oceans previously known as Amansia glomerata (or misidentified as Melanamansia glomerata.13). rhodantha from Western Australia (MURU B262. generally smaller. the Philippines. having tight rosettes as well as very lax. Eastern Australia. Fig. serrate branches and lax ‘rosettes’ of secondary blades issued from the weakly developed midrib. and a cross section of the blade clearly shows the five pericentral cells around the axial ring. From this study. Agardh from Hawaii (LD 42600 / 0772) clearly shows the presence of paired dorsal pseudo-pericentral cells in a blade cross section (Fig. 2) indeed shows a plant with loriform. More importantly. the Cook Islands and French Polynesia (Table I) with A. 3) which distinguishes that plant from A. However. between 1838 and 1842 (US 04057b. The drawing of the holotype (South and Skelton 1999: 246. is comparable to A. The reproductive and trichoblast anatomy of A. with for instance A. one of which (US 04057a) is Amansia rhodantha. paloloensis. isotype and other collections of A. rhodantha (South and Skelton 1999: 249). since the presence of pseudo-pericentral cells is an inconsistent attribute in species of Neurymenia. The present author has re-examined the holotype. paloloensis from the type locality. and found a significant number of thalli with morphologically distinct primary corticated axes and relatively tight and lax rosettes of secondary blades on rosette axes (Figs 11. New Caledonia. Smith (1979: 38–40) points out that the accounting and . While there is no doubt as to the generic identity of the latter specimen. indicating that the morphology of the species is influenced perhaps by environmental factors. loriform blades reminiscent of some specimens of A. D. apparently because the brown pigments (presumably polyphenols) are less susceptible to the bleaching action of the preservative (as is the case in species of brown algae such as Sargassum and Turbinaria). from Melanamansia glomerata was found: when preserved in 4% Formalin-seawater and exposed to ambient light for several days. and Ceramium. paloloensis. 12). 23) but the plant has not been collected in Fiji waters since. Kuetzingia. rhodantha from the type collection and localities in Mauritius. and these authors considered the lack of highly corticated axes. Characters such as the degree of rosette formation. fig. Vidalia. while Melanamansia becomes dark brown. including the presence of both tight and lax rosettes of secondary branches and alar cells on the primary axes. paloloensis showed no major difference in the range of characteristics among them. com. These plants also had blades with well-developed midribs in the lower half (Fig. The Wilkes collection consists of two plants. Fiji. and a weakly developed midrib as characters distinguishing A. Rotuma. rhodantha as described and illustrated by Norris (1988b: 211). figs 1–4) indicate features quite similar to those found in the Amansia species considered in this study from localities in both the Indian Ocean and the tropical Pacific Ocean. N’Yeurt Discussion The holotype of Delesseria rhodantha Harvey An examination by the author of a rehydrated blade fragment of the holotype of Delesseria rhodantha Harvey in TCD (Fig. On the basis of these observations. Japan. Mozambique and Western Australia.
so it cannot be confirmed at this stage if M. in some cases. 13). and is backed by other morphological and biochemical differences (Norris 1988b. Herbarium and liquid-preserved samples of Amansia and Melanamansia species from the Pacific. paloloensis revealed a wide range in the degree of rosette formations and cortication of the secondary axes and presence of alae on primary axes. an examination of the holotype and liquid-preserved and pressed isotypes of A. the vesiculate (in Amansia) or non-vesiculate (in Melanamansia) nature of trichoblasts. These ventral. 1) shows both the presence of tight and lax rosettes of secondary branches. pers. Masuda. Norris (1988b: 211) reported that rosette axes in . Herbarium specimens of ‘Amansia glomerata’ from southern Japan housed in SAP (Hokkaido University) were examined. Melanamansia glomerata is unlikely to be confused with other Melanamansia species because of its unique combination of characteristics (Norris 1988b. which is a characteristic feature of both Amansia and Melanamansia (Norris 1988b) can vary from lax to tight. with the blades smooth and superficially reminiscent of those of some Sargassum species (Figs 22–24). degree of marginal serrations and thallus flattening. There is a possibility it might have been mixed with collections from neighbouring localities such as the Solomon Islands. pl. (2000: 188. Payri and N’Yeurt 1997) were reexamined. they were clearly apparent and about half the size of pericentral cells (Fig. From an examination of the specimens at hand. rhodantha. glomerata occurs in the Solomon Islands. which also had a consistently thicker thallus. while younger plants have less cortication and more lax or absent rosettes of blades. while in Malaysia Masuda et al. All Amansia rhodantha specimens examined had fairly prominent. pseudo-pericentral cells were universally absent. For instance. The author has unsuccessfully tried to obtain material from the latter locality. perhaps an adaptation to the exposed habitats where the plants came from. and found to consistently lack pseudo-pericentral cells and a similar observation was made on samples from Taiwan. Fiji and New Caledonia (Table I). both on the same plant and within individuals in similar populations. paloloensis. and some samples from Kenya. Generally.Revision of some Amansia and Melanamansia species 239 recording of the Wilkes Expedition collections was not very satisfactory. 5. M. position of the reproductive axes and the distinctness of the axial ring in rosette axes (Norris 1988a. L 0108603). fig. Wilson and Kraft 2000). Proliferations from the ventral midrib (aside from secondary branchlets) were only found to be abundant in some of the Indonesian Amansia rhodantha specimens from Salayer (e. 2).g. except the Hawaiian samples. 41) similarly report the absence of pseudo-pericentral cells in A. The illustration by Harvey of the holotype of Delesseria rhodantha (1834. When pseudo-pericentral cells were present. irregularly dichotomous proliferations lack alar cells. in New Caledonia and Kenya). Validity of current criteria to distinguish the Amansieae Characters that have proved useful in distinguishing species in the Amansieae include the presence or absence of pseudo-pericentral cells. Indonesian and Indian Ocean region were sectioned and examined for pseudo-pericentral cells. 7). and found to lack pseudo-pericentral cells. 19). glomerata occurs in the same habitat as Amansia rhodantha (e. However. environmental factors seem to play a role in determining the degree of rosette formation. rhodantha. the Philippines. Fijian and French Polynesian specimens previously ascribed to Melanamansia (N’Yeurt et al. and a main axis with alar cells not very distinct from secondary branches. however it is clear that the degree of rosette formation and cortication of the axes is not a reliable taxonomic character at the species level in Amansia. fig. M. adding to the confusion between the two genera.). the presence or absence of bladelets in rosettes. when present).e. Ecological studies would be needed to verify these hypotheses. com. the description of the species states that branches have the same shape as the main axis (i. in the case of Amansia paloloensis). some specimens were virtually indistinguishable from specimens of A. more exposed conditions (such as on fringing reefs) favour tight rosette formation (which is more hydrodynamically resistant) while more lax arrangements are possible in deeper. rhodantha from Mauritius and Western Australia (compare Figs 4. thus placing them in Amansia. In Melanamansia glomerata these marginal serrations were much less pronounced (but more regular than in Amansia rhodantha. which was the main reason for separating the species from A. fig. more corticated secondary and primary axes with tight rosettes. in the case of A. However. and hence the precise locality and date of this Melanamansia collection from Fiji is unknown. both have an ala) and are arranged in ‘lax’ rosettes (South and Skelton 1999: 247. Indeed. In most instances. calm waters (such as blue holes. Also. older plants tend to have thicker. It would seem that perhaps shallow. and in some instances lacking altogether. 1995). alternate marginal proliferations (which represent non-determinate or endogenous secondary branchlets) 2–5 mm long (Fig.g. Indian Ocean Amansia rhodantha also lacked such cells. and the presence of tight and lax rosettes of secondary branches on the same plant (Figs 11–13). and would seem to function as attachment haptera. it was observed that rosettes of blades along the secondary axes. 1996. The presence or absence of these pseudo-pericentral cells hence appears to be a good generic criterion. CXXVI. type of endogenous branching.
which was not seen in Amansia rhodantha (Figs 6. showing the currently known distribution. This also varied according to the degree of rosette formation and age of the plant. These differences reinforce the conspecificity of the Amansia rhodantha and A. However. as well as more corticated axes. R. At this stage there seems to be no largely significant difference in the number of segments between alternating veins in the specimens of A. it was noted that in Melanamansia the range was more restricted. as well as between A. rhodantha specimens observed in this study. From Table I. There appears to be no notable difference in the range of thallus size. com. paloloensis specimens studied. Fig. rhodantha and A.4 (Norris 1995). but can also occur apically.240 A. These can be easily detected in the unrolled apical portions of the blades by the developing trichoblast terminating each such alternating ‘vein’ (Norris 1995 and pers. smaller and narrower blades. rhodantha studied. usually 3 or 4 with an average of 3.). with the length of segments ranging from 80 to 120 µm. only being more or less surrounded by cortication. 16) and Melanamansia show no particular difference. paloloensis. in elongate flattened and pinnate stichidia which are usually on lateral marginal branchlets. N’Yeurt Amansia from Natal have an ill-defined axial row of cells. An additional potentially useful character is the number of segments between lateral ‘veins’ on the blades. Norris (1995: 66) counted the number of segment cells between these alternating veins for several species of Melanamansia and Amansia. Biogeographical map for Amansia rhodantha (circle) and Melanamansia glomerata (triangle). which was also recorded by Norris (1988b) for Hawaiian material. D. In Melanamansia it averages 4.3. with most specimens having segment lengths of about 100 µm. Cells of the blade alae in both Amansia and Melanamansia are regularly elongate-hexagonal. as well as the generic distinctness of Melanamansia glomerata. blade dimensions. which are in fact determinate branches covered by the pericentral cell derivatives. 15). as the axial ring was still discernible in all sections made. and found them relatively constant. in that they are typically rhodomelaceous with pairs of tetrasporangia in up to ten rows. Tetrasporangial stichidia are basically similar in both Amansia rhodantha and Melanamansia glomerata. but the range is reportedly less than in Amansia rhodantha. 25. An examination of these elegant cells in species of Amansia (Figs 10. In A. cortication of axes and rosette formation between specimens of Amansia rhodantha studied. . Hence the distinctness of the axial ring was not found a reliable taxonomic character among the species examined. a number of preliminary conclusions can be made: A. Cover cells usually totally cover the developing sporangia. This feature was examined on a range of specimens of Amansia from the South Pacific and the Indian Ocean but no significant difference was seen in these specimens. and occur in two overlapping and slightly offset layers radiating on either side of the midrib. 20). Melanamansia glomerata from all localities studied have consistently smoother. the number ranged from 3 to 7. B. One feature which was noted in Melanamansia glomerata is that individual tetrasporangial stichidia are often laterally branched once or twice (Fig.
J. Kylin. et description de cinq nouveaux genres de cette famille. 1956. Hooker by the late Mrs.. V. I. Mus. It is clear that both A. Lewis. Lund). trichoblasts on every second or third axial cell. 458 figs. Nat. – Absence of paired dorsal pericentral cells. Bishop Museum Press. Agardh. A. J.. H. ser. 1963. W. F. 71 pls. and it appears that Melanamansia glomerata is so far restricted to Hawaii (Norris 1995). K. J. Gleerups Förlag.. Cribb. Isaac. Robin South and Mr Posa A. Cambridge University Press. J. Professor G.). xvi + 754 pp. K. Mar. 1901. Sci. 7th edition. composed of up to 4 cells: Melanamansia glomerata. Dr John Parnell (TCD – Trinity College. 1956. and J. Notice of a collection of algae. Bull. Bohn. Bot. Falkenberg. 1841.A. . Washington). Cal. Systema algarum. Mar. Bot. pig- References Abbott. W.. Utrecht. Norris is thanked for his helpful insights and suggestions for this research. In historiam algarum symbolae. 22: 161–193. Honolulu. Professor G. ments rose-red. pigments reddish-brown. Contrib. W. African Bot. frontispiece. Schofield. com. 43: 315–346. Lamouroux. 1968.W. Berlin.A. Dr R. initially vesiculate and composed of up to 8 cells: Amansia rhodantha. Hist. Seven species of Rhodophyceae. Essais sur une classification des algues et des polypiers calcifères de Lamouroux. 1809. 2 folded maps. Mar. 24 pls. J. Leiden). 2. South pers. C. 27: 7–28. M. Charles Telfair. Lord Howe Island (G. Hommersand. Isaac. Nouv. J. Rhodophyta. Linnaea 15: 1–50. Micronesica 21: 53–70. The morphology and classification of some Ceramiaceae and Rhodomelaceae. 2000. Observations sur la physiologie des algues marines. 443–457. Millar. E.Vietnam and Malaysia (M. Mr Barrett Brooks and Dr John Sims (US – Smithsonian Institution. 17: 297–380. H. 1981. C. The Herbaria of the World. S. (2000) confirm that the genus Amansia does exist in the Pacific Ocean. A. Australian Coral Reef Society: Brisbane. Paris 1: 330–333. from “Cap Malheureux”. S. Phil. Index Herbariorum. W. J. Isaac. II. with descriptions of some new and little known species. Jaasund. Garrigue. First edition. Marine red algae from the North coast of Papua New Guinea. E. 336 figs. (Hooker) 1: 147–157. flora and vegetation. Fauna und Flora des Golfes von Neapel. A history and annotated account of the benthic marine algae of Taiwan. M. U. Marine Red Algae of the Hawaiian Islands. Sapporo). G. Smiths. 10 figs. Coppejans. E.. and F. Masuda pers. Catalogue of marine benthic algae from New Caledonia. Sci. Marine Algae of the Southern Great Barrier Reef. Accepted 17 January 2002. and A. Sci. M. 1983. pls cxxv. Bot. trichoblasts non-vesiculate. Soc. T. Phang.. Holmgren. F. 29: 1–38. 2000. Agardh. 173 + (2) pp. Volume II. E. 43: 181–190. R. Marine algae of Inhaca Island and of the Inhaca Peninsula. Part 1. Lunda (Lund). Millar for his generous assistance in obtaining copies of rare publications. 1945. B. Dublin). Skelton (SUVA – The University of the South Pacific). 1987. 1988. xvi + 477 +  pp. E. Bot. University of Tromsø. General account of the environment. Decaisne. Masuda. S. Acknowledgements The author sincerely thanks the following persons and institutions for the generous loan of type material and specimens used in this study: Dr John Huisman (MURU – Murdoch University). C. New Caledonia and the Philippines (this study). communicated to Dr. K. Phaeophyceae. Scheltema & Holkema. xv + 673 pp. Revised Key to the Species of Amansia and Melanamansia in the South Pacific – Presence of paired dorsal pericentral cells. East Africa Nat. Rhodophyceae. Foreword. com. xiv + 939 pp. sparse. Monographie 26. Die Rhodomelaceen des Golfes von Neapel und der angrenzenden Meeres-Abschnitte. Soc. Taxonomic notes on marine algae from Malaysia. Norris.. iii + 159 pp. The Structure and Reproduction of the Algae. Kato. 1824. Dr Claude Payri (UPF – Tahiti).). E. Nat. Pub.Revision of some Amansia and Melanamansia species 241 Conclusions The results of this study and those of South and Skelton (1999) and of Masuda et al. rhodantha and Melanamansia glomerata are more widely distributed than previously thought. J. Harvey. Professors Michio Masuda and Tadao Yoshida (SAP – Hokkaido University. 1834. P. K. Bot. Handbook No 2. Keuken and E. Kenya (this study). and each genus is consistently differentiated from the other by a number of anatomical and biochemical characters. 1976. N. Ms Susanna Riebe (LD – Botanical Museum. and Dr A. E. M. Shimada. Die Gattungen der Rhodophyceen. Lund. and R. P. xxxviii + 312 pp. H. 1842. in the Mauritius. Tsuda. Figure 25 presents a world distribution of these two genera as it is currently known. Part I. pls v–vii. Kawaguchi and S. K. Myxophyceae. 35: 165–366. and possibly other localities which are not yet adequately investigated. Marine botany of the Kenya coast. cxxvi. Annal. 1999. Professor Willem Prud’homme van Reine (L – Rijksherbarium. 3. Intertidal Seaweeds in Tanzania. Fritsch. I. Robin South is also thanked for his helpful comments on an earlier draft of this manuscript. J.
Kauai. G. Lawai. Skelton. R. Publ. Catalogue of the benthic marine algae of the Indian Ocean. 1997.. U. Vol. Austr. Micronesica 29: 49–96. T. A revised checklist of Polynesian benthic marine algae. pp. C. 27 May–1 June 1985. P. 1996. Calif. N’Yeurt. 1999. v. and A. and A. and P. Illustrations of the Marine Algae of Japan I. J. Systematic Bot. figs 110–142. Galzin and B. Algol. nov. Bot. Silva. Jap. D. 1. 24: 209–223. C. Liste des Algues du Siboga. French Polynesian Coral Reefs. Pap. Norris. (Rhodomelaceae. J. 2000. Yamada. E. S. Volume 1. Flora Vitiensis Nova. 1923. Keats. Seconde Partie. Micronesica 13: 85–120. Wray. 498–518... D. Payri. South. A revised checklist of the benthic marine algae of Fiji (including the island of Rotuma).A. and T. comb. K. Melanamansia glomerata. E. J. R. C. Salvat. In: (B. (Sôrui) 43: 115–171. eds) Proceedings of the 5th International Coral Reef Congress. Algae. Hokkaido Imperial U. Bibliography of marine benthic algae in Micronesia. Y. Phycol. Payri. Amansia paloloensis sp. Rhodophyta. South Pacific. 1900–1902.. Austr. 1979. Inst. Tokyo. M. Tanaka. T. Wilson. 1996. Nakajima. Contrib. Antenne de Tahiti. P. 1995.. R. 1996. Basson and R. R. A. 1–93. 12: 549–591. Keigyosha and Co. 9: 361–490. R.. A. and Amansia rhodantha. G. C. D.. A. J. Rhodomelaceae) on the southeastern African coast. Morphological and taxonomic studies on selected genera from the tribe Amansieae (Rhodomelaceae.. The marine algae from the island of Yonakuni. Rhodophyta) from Samoa. Hawaii. Austr. Moorea. R. Systematic Bot. A New Flora of Fiji. K. 311–392. 27: iv + 1–179. W. Yoshinaga and Y. Sci. Papua New Guinea). Sci. 1999. Smith. Okamura. Moe. pp. Millar. Taxon 44: 65–68. Norris. pp. E. 1985. 12: 479– 547. xxx pls. 1987. and G. Meñez and R. M. Smiths. C. R. A. Weber-van Bosse. 1999. L. Marine benthic algae of Norfolk Island. Coppejans and L. pls ix–x. E. N’Yeurt. D. R. Phycol. O. L. III. 79: 1–1259. J. Fac.242 A. Tahiti. Delesalle. P. A preliminary floristic survey of the benthic marine algae of Rotuma Island. Tsuda. Rhodophyta). Rhodomelaceae) in southeastern Africa. (Sôrui) 36: 271–276. III. 10: 867–910. K. G. Ceramiales. R. Moe. 1995. (Rhodophyta.. Liao. Jap. Rhodophyceae. N’Yeurt. Austr. A. Annotated and illustrated survey of the marine macroalgae from Motupore Island and vicinity (Port Moresby area. nov. Meinesz. E. nov. W. South Pacific. Norris. Catalog of the Marine Benthic Algae of the Philippines. Austr. The specific identity of Neurymenia (Rhodophyceae. 13: 325–372. Mar. Phycol. Res. R. . E. Muséum National d’Histoire Naturelle. Kraft. Systematic Bot. Sci. 1977. 1988a. and F. E. Phycologia 38: 245–250. A. Silva. N’Yeurt Millar. École Pratique des Hautes Études. Siboga-Expeditie Monographie 59c. ill. 2: 53–86. Systematic Bot. de Clerk. 1988b. 1938. Structure and reproduction of Amansia and Melanamansia gen. Yoshida. South and D. R. O.. two hitherto confused species of Indo-Pacific Rhodophyceae. Checklist of marine algae of Japan (revised in 1995). K. Systematic Bot. T. pp. Pacific Tropical Botanical Garden.
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